Genetic Diversity of Three European Veratrum Species Revealed by Amplified Fragment Length Polymorphism

Total Page:16

File Type:pdf, Size:1020Kb

Genetic Diversity of Three European Veratrum Species Revealed by Amplified Fragment Length Polymorphism Biodiv. Res. Conserv. 47: 1-8, 2017 BRC www.brc.amu.edu.pl DOI 10.1515/biorc-2017-0011 Submitted 13.06.2017, Accepted 30.09.2017 Genetic diversity of three European Veratrum species revealed by Amplified Fragment Length Polymorphism Magdalena Szeliga*, Joanna Ciura & Mirosław Tyrka Department of Biotechnology and Bioinformatics, Faculty of Chemistry, Rzeszow University of Technology, Powstańców Warszawy 6, 35-959 Rzeszów, Poland *corresponding author (e-mail: [email protected]) Abstract. Chemical and genetic characterization of Veratrum species deposited in European collections is important for genepool preservation and identification of populations with desired metabolic properties.Veratrum album, V. lobelianum and V. nigrum are native to Europe, and in Poland are ranked as rare or threatened. Genetic variation of European Veratrum species was characterized by Amplified Fragment Length Polymorphism (AFLP) markers. The accumulation of jervine as a represen- tative of steroidal alkaloids was measured in seeds. Distribution of 380 markers generated from eight primer combinations was useful for studying genetic relationships among and within species in the Veratrum genus and the most divergent populations were identified. Genetic variation between 12 populations ofVeratrum species supports the classification ofV. lobelianum as a subspecies of V. album. However, the results need further validation on extended material. A higher genetic diversity (22.3%) was observed between populations of V. nigrum as compared to V. album (14.5%). Contents of jervine allowed for discrimina- tion of the studied Veratrum species and can be used as a potential chemotaxonomic marker. The highest jervine levels were found in V. album. V. nigrum seeds had only trace amounts and no jervine was detected in seeds of V. lobelianum. Key words: genetic variation, hellebore, jervine, Melanthiaceae, molecular markers 1. Introduction are divided according to structural features (Li et al. 2006). These steroidal alkaloids are well known for The Veratrum genus (Liliales, Melanthiaceae) com- their pharmacological activities, including hypotensive, prises 17-45 species distributed over a wide range of antithrombotic and antitumour functions (Tang et al. habitats of the Northern Hemisphere (Zomlefer et al. 2008a; Ivanova et al. 2011). In particular, cyclopamine 2003; Treier & Müller-Schärer 2011). Veratrum album showed antitumour activity and induced apoptosis using L., Veratrum lobelianum Bernh. and Veratrum nigrum the mechanism of inhibition of the hedgehog pathway in L. are native to Europe. Only a few isolated populations a subset of the pancreatic cancer cell line (Thayer et al. of these species are present in Poland and ranked as rare 2003). Due to low cyclopamine concentration in Vera- or threatened (Allen et al. 2014). Veratrum album sensu trum plants (about 0.01%), we focussed our attention on lato is a complex of subspecies V. album ssp. album jervine that was present in higher concentrations (0.1%) and V. album ssp. lobelianum (Schaffner et al. 2001) and can be transformed to cyclopamine by Wolff-Kish- sometimes considered as separate species (Zomlefer et ner reduction with Huang-Minlon modification (Tanget al. 2003). Plants of Veratrum are a rich source of unique al. 2008b). Jervine displays strong antifungal activities bioactive steroidal alkaloids (Chandler & McDougal against the phytopathogenic Phytophthora capsici (Li 2014). Quantitative and qualitative variations in the et al. 2006). metabolic profile and pharmacological properties are Medical plants from Melanthiaceae are interest- affected by environmental factors and genetic diversity ing specimens for genetic diversity studies (Li et al. present in natural populations (Li et al. 2015). 2011; Treier & Müller-Schärer 2011). For genetic There are three subtypes of Veratrum-type alka- characteristics targeting taxonomic questions, variation loids including cevanine, veratramine and jervine that in cpDNA and the internal transcribed spacer (ITS) AND PHYLOGENY TAXONOMY VARIABILITY, ©Adam Mickiewicz University in Poznań (Poland), Department of Plant Taxonomy. All rights reserved. 2 Magdalena Szeliga et al. Genetic diversity of three European Veratrum species revealed by amplified fragment... region of rDNA are standard (Zomlefer et al. 2003; 2. Materials and methods Griffin & Barrett 2004; Liao et al. 2007). Systems targeting conservative regions of DNA were ideal 2.1. Plant materials for separating evolutionary different populations of Trillium grandiflorum and V. album ssp. oxysepalum Plant collection consisted of six populations of (Griffin & Barrett 2004; Kikuchiet al. 2010). However, V. nigrum, five populations of V. album and a single to study genetic diversity within and between natural population of V. lobelianum maintained in botanical gar- populations, amplified fragment length polymorphism dens across seven European countries (Table 1). Seeds (AFLP) system is a method of choice (Guthridge et al. from population N5 (V. nigrum) were germinated and 2001; Quagliaro et al. 2001; Tang et al. 2003; Llanes plants were maintained in vitro, then lyophilized prior et al. 2011). This DNA fingerprinting technique targets to DNA extraction. Seeds of the remaining families had multiple loci, generates dominant markers for whole- reduced viability and, therefore, were used directly (3-7 genome screening and has higher reproducibility, reso- random seeds) for DNA isolation. The names of genus lution, and sensitivity compared to random amplified and species follow Zomlefer et al. (2003). polymorphic DNA (RAPD) and inter simple sequence 2.2. DNA extraction and AFLP analysis repeats (ISSRs) methods (Vos et al. 1995; Blears et al. 1998). Total plant genomic DNA was extracted from seeds Various genetic marker systems have been used to or lyophilized plants using a method developed by Mil- characterize variation in the Veratrum genus, including ligan (1992). The DNA was quantified using agarose RAPD (Kleijn & Steinger 2002), AFLP (Treier & Mül- gel electrophoresis and diluted to a concentration of ler-Schärer 2011) and simple sequence repeats (Kato et 200 ng·mL-1 prior to AFLP analysis. The AFLP analysis al. 2008; Kikuchi & Maki 2011). AFLP studies of 40 was carried out according to the methods of Thomas et European natural populations of V. album revealed east- al. (1995). Genomic DNA was digested with MseI and west direction in the genetic structure and suggested PstI restriction enzymes (5 U each) in Tango buffer (33 ancient migration from Asia (Treier & Müller-Schärer mM Tris-acetate, pH 7.9 at 37°C; 10 mM magnesium 2011). acetate; 66 mM potassium acetate; 0.1 mg/ml BSA) for So far, genetic diversity between and within European 90 min at 65°C and 60 min at 37°C, respectively. The accessions of V. nigrum, V. album and V. lobelianum has enzymes were subsequently heat inactivated at 80°C not been investigated with AFLP markers. Chemical and for 20 min and the products ligated with MseI and PstI genetic characterization of Veratrum species deposited adapters overnight at 37°C to generate template DNA in European collections generates new information for amplification. DNA was then purified by ethanol pre- useful both for genepool preservation and for identifica- cipitation and dissolved in 20 µL of water. Non-selective tion of populations with desired metabolic properties. (NS) amplification (30 cycles of 30 s at 94°C, 30 s at Jervine content was determined to preliminary test the 56°C and 60 s at 72°C) was conducted using primers suitability of this genera-specific steroid alkaloid as a complementary to MseI and PstI adaptors without potential chemotaxonomic marker. selective nucleotides (5'-GATGAGTCCTGAGTAA-3' Table 1. Populations of Veratrum species used for genetic and chemical analyses Number Acronym Species Sampling locality Source of material of plants N1 V. nigrum Nowy Sącz, POL Private garden 3 N2 V. nigrum Ljubljana, SVN University Botanic Gardens 7 N3 V. nigrum Bayreuth, DEU Ecological Botanical Gardens 5 N4 V. nigrum Romsey, GBR Sir Harold Hillier Gardens 5 N5 V. nigrum Brno, CZE Masaryk University Botanic Garden 6 N6 V. nigrum Yoskar-Ola, RUS Botanical Garden of the Technological State University 5 L1 V. lobelianum Ljubljana, SVN University Botanic Gardens 6 A1 V. album Bayreuth, DEU Ecological Botanical Gardens 5 A2 V. album Bolestraszyce, POL Arboretum and Institute of Physiography 6 A3 V. album Bonn, DEU University Botanic Gardens 6 A4 V. album Nancy, FRA Conservatory and Botanical Gardens 4 A5 V. album High Tatra Mts, POL Collecting 6 Biodiv. Res. Conserv. 47: 1-8, 2017 3 and 5'-GACTGCGTACATGCAG-3', respectively). following transitions were detected: m/z 425.89/313.10 The diluted PCR products of the NS amplification were Da and 425.89/114.0 Da for jervine (Tang et al. 2008b; selectively amplified with eight combinations of MseI Grobosch et al. 2008). primers (M43, M47, M48, M50, M59) and PstI primers 2.4. Data analysis (P4, P12, P15, P16, P18, P26, P34), each containing three selective nucleotides at the 3'-end. Symbols of the For the diversity analysis, the amplification products selective primers are in accordance with the KeyGene were scored for the presence (1) and absence (0) of standard list for AFLP primer nomenclature. Selective bands to form a binary matrix. These data were used amplification was accomplished with a touchdown to calculate polymorphism information, including the thermal cycle, as follows: 7 cycles at 94°C for 60 s, 65°C number of polymorphic/species-specific fragments
Recommended publications
  • Leprosy and Other Skin Disorders
    Copyright by Robert Joseph Gallagher 2014 The report committee for Robert Joseph Gallagher Certifies that this is the approved version of the following report: An Annotated Translation of Chapter 7 of the Carakasaṃhitā Cikitsāsthāna: Leprosy and Other Skin Disorders APPROVED BY SUPERVISING COMMITTEE: Supervisor: __________________________________ Donald R. Davis _________________________________ Joel Brereton An Annotated Translation of Chapter 7 of the Carakasaṃhitā Cikitsāsthāna: Leprosy and Other Skin Disorders by Robert Joseph Gallagher, B.A., M.A. Report Presented to the Faculty of the Graduate School of The University of Texas at Austin in Partial Fulfillment for the degree of Master of Arts University of Texas at Austin May 2014 Dedication To my wife Virginia and our two daughters Michelle and Amy, who showed patience and understanding during my long hours of absence from their lives, while I worked on mastering the intricacies of the complex but very rewarding language of Sanskrit. In addition, extra kudos are in order for thirteen year-old Michelle for her technical support in preparing this report. Acknowledgements I wish to thank all the members of the South Asia team at UT Austin, including Prof. Joel Brereton, Merry Burlingham, Prof. Don Davis, Prof. Oliver Freiberger, Prof. Edeltraud Harzer, Prof. Patrick Olivelle, Mary Rader, Prof. Martha Selby and Jennifer Tipton. Each one has helped me along this path to completion of the M.A. degree. At the time of my last serious academic research, I used a typewriter to put my thoughts on paper. The transition from white-out to pdf has been challenging for me at times, and I appreciate all the help given to me by the members of the South Asia team.
    [Show full text]
  • A Medical-Historical Examination of the Death of Alexander the Great
    A MEDICAL-HISTORICAL EXAMINATION OF THE DEATH OF ALEXANDER THE GREAT Abstract: Alexander the Great’s cause of death has been contentious since antiquity. Historians and physicians alike have proposed a multitude of hypotheses. However, neither party is without their analytical flaws. The historians often neglect obvious medical refutations. Meanwhile, the physicians often err by forsaking disciplined historical methodology. Therefore, the authors of this paper subject these prior hypotheses to both Nathan Gamble medical and historical criticism, in order to provide a multidisciplinary University of Toronto, Toronto, Canada approach to a longstanding mystery. Some hypotheses have more weight St. Mary’s University, Twickenham, London than others, as is discussed. The most probable of the poisoning hypotheses, [email protected] which aligns with the Vulgate tradition of Alexander’s death, cites the use of Veratrum album, a plant derived bane. When the Court tradition is considered, Edmund F. Bloedow i.e. that no foul play occurred, acute pancreatitis induced by alcohol abuse holds greatest credence as a hypothesis. It is hoped that the approach used Augustine College, Ottawa, Ontario will not only increase clarity regarding Alexander’s death and challenge weak [email protected] ideas but also provide an approach by which speculation about other medical diagnoses in history may be tempered and critiqued. Keywords: Alexander the Great, Retrospective Diagnosis, Regicide, Murder, Death DOI: 10.14795/j.v4i3.269 ISSN 2360 – 266X ISSN–L 2360 – 266X ACKNOWLEDGMENTS: The authors would like to thank Joel Gamble and Jonathan Madany for the kind help in reviewing and preparing the manuscript for publication.
    [Show full text]
  • Liliaceae S.L. (Lily Family)
    Liliaceae s.l. (Lily family) Photo: Ben Legler Photo: Hannah Marx Photo: Hannah Marx Lilium columbianum Xerophyllum tenax Trillium ovatum Liliaceae s.l. (Lily family) Photo: Yaowu Yuan Fritillaria lanceolata Ref.1 Textbook DVD KRR&DLN Erythronium americanum Allium vineale Liliaceae s.l. (Lily family) Herbs; Ref.2 Stems often modified as underground rhizomes, corms, or bulbs; Flowers actinomorphic; 3 sepals and 3 petals or 6 tepals, 6 stamens, 3 carpels, ovary superior (or inferior). Tulipa gesneriana Liliaceae s.l. (Lily family) “Liliaceae” s.l. (sensu lato: “in the broad sense”) - Lily family; 288 genera/4950 species, including Lilium, Allium, Trillium, Tulipa; This family is treated in a very broad sense in this class, as in the Flora of the Pacific Northwest. The “Liliaceae” s.l. taught in this class is not monophyletic. It is apparent now that the family should be treated in a narrower sense and some of the members should form their own families. Judd et al. recognize 15+ families: Agavaceae, Alliaceae, Amarylidaceae, Asparagaceae, Asphodelaceae, Colchicaceae, Dracaenaceae (Nolinaceae), Hyacinthaceae, Liliaceae, Melanthiaceae, Ruscaceae, Smilacaceae, Themidaceae, Trilliaceae, Uvulariaceae and more!!! (see web reading “Consider the Lilies”) Iridaceae (Iris family) Photo: Hannah Marx Photo: Hannah Marx Iris pseudacorus Iridaceae (Iris family) Photo: Yaowu Yuan Photo: Yaowu Yuan Sisyrinchium douglasii Sisyrinchium sp. Iridaceae (Iris family) Iridaceae - 78 genera/1750 species, Including Iris, Gladiolus, Sisyrinchium. Herbs, aquatic or terrestrial; Underground stems as rhizomes, bulbs, or corms; Leaves alternate, 2-ranked and equitant Ref.3 (oriented edgewise to the stem; Gladiolus italicus Flowers actinomorphic or zygomorphic; 3 sepals and 3 petals or 6 tepals; Stamens 3; Ovary of 3 fused carpels, inferior.
    [Show full text]
  • Guide to the Flora of the Carolinas, Virginia, and Georgia, Working Draft of 17 March 2004 -- LILIACEAE
    Guide to the Flora of the Carolinas, Virginia, and Georgia, Working Draft of 17 March 2004 -- LILIACEAE LILIACEAE de Jussieu 1789 (Lily Family) (also see AGAVACEAE, ALLIACEAE, ALSTROEMERIACEAE, AMARYLLIDACEAE, ASPARAGACEAE, COLCHICACEAE, HEMEROCALLIDACEAE, HOSTACEAE, HYACINTHACEAE, HYPOXIDACEAE, MELANTHIACEAE, NARTHECIACEAE, RUSCACEAE, SMILACACEAE, THEMIDACEAE, TOFIELDIACEAE) As here interpreted narrowly, the Liliaceae constitutes about 11 genera and 550 species, of the Northern Hemisphere. There has been much recent investigation and re-interpretation of evidence regarding the upper-level taxonomy of the Liliales, with strong suggestions that the broad Liliaceae recognized by Cronquist (1981) is artificial and polyphyletic. Cronquist (1993) himself concurs, at least to a degree: "we still await a comprehensive reorganization of the lilies into several families more comparable to other recognized families of angiosperms." Dahlgren & Clifford (1982) and Dahlgren, Clifford, & Yeo (1985) synthesized an early phase in the modern revolution of monocot taxonomy. Since then, additional research, especially molecular (Duvall et al. 1993, Chase et al. 1993, Bogler & Simpson 1995, and many others), has strongly validated the general lines (and many details) of Dahlgren's arrangement. The most recent synthesis (Kubitzki 1998a) is followed as the basis for familial and generic taxonomy of the lilies and their relatives (see summary below). References: Angiosperm Phylogeny Group (1998, 2003); Tamura in Kubitzki (1998a). Our “liliaceous” genera (members of orders placed in the Lilianae) are therefore divided as shown below, largely following Kubitzki (1998a) and some more recent molecular analyses. ALISMATALES TOFIELDIACEAE: Pleea, Tofieldia. LILIALES ALSTROEMERIACEAE: Alstroemeria COLCHICACEAE: Colchicum, Uvularia. LILIACEAE: Clintonia, Erythronium, Lilium, Medeola, Prosartes, Streptopus, Tricyrtis, Tulipa. MELANTHIACEAE: Amianthium, Anticlea, Chamaelirium, Helonias, Melanthium, Schoenocaulon, Stenanthium, Veratrum, Toxicoscordion, Trillium, Xerophyllum, Zigadenus.
    [Show full text]
  • Cally Plant List a ACIPHYLLA Horrida
    Cally Plant List A ACIPHYLLA horrida ACONITUM albo-violaceum albiflorum ABELIOPHYLLUM distichum ACONITUM cultivar ABUTILON vitifolium ‘Album’ ACONITUM pubiceps ‘Blue Form’ ACAENA magellanica ACONITUM pubiceps ‘White Form’ ACAENA species ACONITUM ‘Spark’s Variety’ ACAENA microphylla ‘Kupferteppich’ ACONITUM cammarum ‘Bicolor’ ACANTHUS mollis Latifolius ACONITUM cammarum ‘Franz Marc’ ACANTHUS spinosus Spinosissimus ACONITUM lycoctonum vulparia ACANTHUS ‘Summer Beauty’ ACONITUM variegatum ACANTHUS dioscoridis perringii ACONITUM alboviolaceum ACANTHUS dioscoridis ACONITUM lycoctonum neapolitanum ACANTHUS spinosus ACONITUM paniculatum ACANTHUS hungaricus ACONITUM species ex. China (Ron 291) ACANTHUS mollis ‘Long Spike’ ACONITUM japonicum ACANTHUS mollis free-flowering ACONITUM species Ex. Japan ACANTHUS mollis ‘Turkish Form’ ACONITUM episcopale ACANTHUS mollis ‘Hollard’s Gold’ ACONITUM ex. Russia ACANTHUS syriacus ACONITUM carmichaelii ‘Spätlese’ ACER japonicum ‘Aconitifolium’ ACONITUM yezoense ACER palmatum ‘Filigree’ ACONITUM carmichaelii ‘Barker’s Variety’ ACHILLEA grandifolia ACONITUM ‘Newry Blue’ ACHILLEA ptarmica ‘Perry’s White’ ACONITUM napellus ‘Bergfürst’ ACHILLEA clypeolata ACONITUM unciniatum ACIPHYLLA monroi ACONITUM napellus ‘Blue Valley’ ACIPHYLLA squarrosa ACONITUM lycoctonum ‘Russian Yellow’ ACIPHYLLA subflabellata ACONITUM japonicum subcuneatum ACONITUM meta-japonicum ADENOPHORA aurita ACONITUM napellus ‘Carneum’ ADIANTUM aleuticum ‘Japonicum’ ACONITUM arcuatum B&SWJ 774 ADIANTUM aleuticum ‘Miss Sharples’ ACORUS calamus ‘Argenteostriatus’
    [Show full text]
  • Paton, Anne Elizabeth (2015) an Examination of the Evidence for the Existence of Leprosy and Hansen’S Disease in Medieval Ireland
    Paton, Anne Elizabeth (2015) An examination of the evidence for the existence of leprosy and Hansen’s disease in medieval Ireland. PhD thesis. https://theses.gla.ac.uk/6427/ Copyright and moral rights for this work are retained by the author A copy can be downloaded for personal non-commercial research or study, without prior permission or charge This work cannot be reproduced or quoted extensively from without first obtaining permission in writing from the author The content must not be changed in any way or sold commercially in any format or medium without the formal permission of the author When referring to this work, full bibliographic details including the author, title, awarding institution and date of the thesis must be given Enlighten: Theses https://theses.gla.ac.uk/ [email protected] AN EXAMINATION OF THE EVIDENCE FOR THE EXISTENCE OF LEPROSY AND HANSEN’S DISEASE IN MEDIEVAL IRELAND ANNE ELIZABETH PATON M.A. (Hons.), M.Sc. Submitted in fulfillment of the requirements for the Degree of Ph.D. School of Humanities College of Arts University of Glasgow August 2014 Abstract Much concerning the disease termed leprosy is accepted as received knowledge, without thought to time and place, but there were many differences in how leprosy sufferers were treated across regions and eras, and so diversity should be regarded as normal. This thesis will examine what was meant by the term leprosy during primarily the medieval period between the sixth and fifteenth centuries in Ireland in order to see if this equates with the disease called Hansen’s Disease in the twenty-first century.
    [Show full text]
  • Aconitum Napellus)
    Phil Rasmussen (M.Pharm., M.P.S., Dip. Herb. Med., M.N.H.A.A., M.N.I.M.H.(U.K.), M.N.Z.A.M.H.) Consultant Medical Herbalist 23 Covil Ave Te Atatu South Auckland New Zealand tel.(0064)09 378 9274 fax.(0064) 09 834 8870 email: [email protected] _____________________________________________________________________ Report on Appropriate Classification for Aconite (Aconitum napellus) Confidential May 9, 2001. Summary An assessment of safety considerations with respect to human usage of complementary medicine preparations made from the substance Aconite (any part of the plant Aconitum napella, otherwise known as Monkshood), has been undertaken. The available toxicological data was reviewed, and levels of intake of the known toxic constituents, the alkaloids aconitine, mesaconitine and jesaconitine, known to be associated with adverse effects and possible fatality in humans, were determined. From this assessment, concentration levels of the known toxic alkaloids below which no toxic effects would normally be associated with their internal ingestion or use, was determined. Levels of ingestion of these toxic components which could normally be deemed as completely safe, were then ascertained. This assessment was then applied to an evaluation of homoeopathic Aconite-containing preparations available in the marketplace, to select ‘cut off points’ below which general sales classification is deemed appropriate. These calculations were based upon both concentration levels of the toxic alkaloids, as well as the maximum recommended pack size of preparations containing them. Aconite: an introduction Aconite (a preparation made from either the roots or herb of the European shrub Aconitum napellus, or other Aconitum species ), has long been used both as a traditional herbal medicine as well as a homoeopathic remedy.
    [Show full text]
  • Veratrum Steroidal Alkaloid Toxicity Following Ingestion of Foraged
    Veratrum Steroidal Alkaloid Toxicity Following Ingestion of Foraged Veratrum Parviflorum M Anwar 1, MW Turner 2, N Farrell 3, R Kleiman 4, WB Zomlefer 5, OM McDougal 2, BW Morgan 1 1Emory University School of Medicine, Atlanta, GA; 5Boise State University, Boise, ID; 3Rhode Island Hospital, Providence, RI; 4Wellstar Kennestone Hospital, Marietta, GA; 5University of Georgia, Athens, GA BACKGROUND RESULTS DISCUSSION • Steroidal alkaloids are found in the Veratrum genus of • The specimen was identified as V. parviflorum by botanists • Steroidal alkaloids have previously been isolated and toxicity plants. at the University of Georgia . has been reported from many species of Veratrum plants. ¡ ¢ £ ¡ ¤ ¥ ¢ ¦ ¨ § • Their toxicity manifests as GI illness followed by a x107 1.0 • This is the first reported case of Veratrum toxicity from V. Bezold-Jarisch reflex: hypopnea, hypotension and 0.5 ©¨ parviflorum with identified steroidal alkaloids. bradycardia. x107 1.0 • As far as we know, there is no previous study to characterize • Some Veratrum steroidal alkaloids are also teratogens 0.5 ¨ x107 the steroidal alkaloids in V. parviflorum . interfering with the hedgehog-2 signaling pathway which 1.0 causes cyclopsia and holoprosencephaly. 0.5 ¨ • A prior study shows some cross reactivity between Veratrum x107 1.0 steroidal alkaloids and the digoxin assay but no digoxin CASE PRESENTATION 0.5 ¨ immune fab binding. x107 • A 27 year old man (patient 1) and his 25 year old wife 1.0 0.5 (patient 2) presented to the ED with nausea and 0 x107 f) vomiting after foraging and ingesting what they believed 1.0 to be wild leeks from the Appalachian Trail in Georgia, 0.5 10 USA.
    [Show full text]
  • TELOPEA Publication Date: 13 October 1983 Til
    Volume 2(4): 425–452 TELOPEA Publication Date: 13 October 1983 Til. Ro)'al BOTANIC GARDENS dx.doi.org/10.7751/telopea19834408 Journal of Plant Systematics 6 DOPII(liPi Tmst plantnet.rbgsyd.nsw.gov.au/Telopea • escholarship.usyd.edu.au/journals/index.php/TEL· ISSN 0312-9764 (Print) • ISSN 2200-4025 (Online) Telopea 2(4): 425-452, Fig. 1 (1983) 425 CURRENT ANATOMICAL RESEARCH IN LILIACEAE, AMARYLLIDACEAE AND IRIDACEAE* D.F. CUTLER AND MARY GREGORY (Accepted for publication 20.9.1982) ABSTRACT Cutler, D.F. and Gregory, Mary (Jodrell(Jodrel/ Laboratory, Royal Botanic Gardens, Kew, Richmond, Surrey, England) 1983. Current anatomical research in Liliaceae, Amaryllidaceae and Iridaceae. Telopea 2(4): 425-452, Fig.1-An annotated bibliography is presented covering literature over the period 1968 to date. Recent research is described and areas of future work are discussed. INTRODUCTION In this article, the literature for the past twelve or so years is recorded on the anatomy of Liliaceae, AmarylIidaceae and Iridaceae and the smaller, related families, Alliaceae, Haemodoraceae, Hypoxidaceae, Ruscaceae, Smilacaceae and Trilliaceae. Subjects covered range from embryology, vegetative and floral anatomy to seed anatomy. A format is used in which references are arranged alphabetically, numbered and annotated, so that the reader can rapidly obtain an idea of the range and contents of papers on subjects of particular interest to him. The main research trends have been identified, classified, and check lists compiled for the major headings. Current systematic anatomy on the 'Anatomy of the Monocotyledons' series is reported. Comment is made on areas of research which might prove to be of future significance.
    [Show full text]
  • Liliaceae Lily Family
    Liliaceae lily family While there is much compelling evidence available to divide this polyphyletic family into as many as 25 families, the older classification sensu Cronquist is retained here. Page | 1222 Many are familiar as garden ornamentals and food plants such as onion, garlic, tulip and lily. The flowers are showy and mostly regular, three-merous and with a superior ovary. Key to genera A. Leaves mostly basal. B B. Flowers orange; 8–11cm long. Hemerocallis bb. Flowers not orange, much smaller. C C. Flowers solitary. Erythronium cc. Flowers several to many. D D. Leaves linear, or, absent at flowering time. E E. Flowers in an umbel, terminal, numerous; leaves Allium absent. ee. Flowers in an open cluster, or dense raceme. F F. Leaves with white stripe on midrib; flowers Ornithogalum white, 2–8 on long peduncles. ff. Leaves green; flowers greenish, in dense Triantha racemes on very short peduncles. dd. Leaves oval to elliptic, present at flowering. G G. Flowers in an umbel, 3–6, yellow. Clintonia gg. Flowers in a one-sided raceme, white. Convallaria aa. Leaves mostly cauline. H H. Leaves in one or more whorls. I I. Leaves in numerous whorls; flowers >4cm in diameter. Lilium ii. Leaves in 1–2 whorls; flowers much smaller. J J. Leaves 3 in a single whorl; flowers white or purple. Trillium jj. Leaves in 2 whorls, or 5–9 leaves; flowers yellow, small. Medeola hh. Leaves alternate. K K. Flowers numerous in a terminal inflorescence. L L. Plants delicate, glabrous; leaves 1–2 petiolate. Maianthemum ll. Plant coarse, robust; stems pubescent; leaves many, clasping Veratrum stem.
    [Show full text]
  • 4. VERATRUM Linnaeus, Sp. Pl. 2: 1044. 1753
    Flora of China 24: 82–85. 2000. 4. VERATRUM Linnaeus, Sp. Pl. 2: 1044. 1753. 藜芦属 li lu shu Chen Xinqi (陈心启 Chen Sing-chi); Hiroshi Takahashi1 Herbs perennial, usually andropolygamous, with short, thick rhizomes and stout, slightly fleshy roots. Stems erect, terete, stout, usually pubescent, leafy, usually enclosed basally by fibers or reticulate fibers formed from disintegrated sheaths. Leaves alternate, sheathed and clasping, broad to narrow, strongly plicately veined, basally usually narrowed. Inflorescence usually a terminal panicle, many flowered. Flowers shortly pedicellate or subsessile, white, yellowish, green, or dark purple-brown, funnelform, cupular to opening flat. Tepals 6, usually free, spreading, persistent in fruit. Stamens 6, inserted at base of tepals; anthers reniform to cordate- orbicular, with locules confluent and dehiscent by an apical valve. Ovary 3-loculed, slightly 3-lobed apically, ovules usually many. Styles 3, short, persistent, stigmatic adaxially. Fruit a septicidal capsule. Seeds several per valve, flattened, narrowly winged. About 40 species: mainly in temperate regions of the N hemisphere; 13 species (eight endemic) in China. 1a. Basal sheath of stem with only longitudinal veins, becoming fibrous when disintegrated. 2a. Leaves glabrous or sparsely pubescent .............................................................................................................. 2. V. oxysepalum 2b. Leaves densely pubescent abaxially. 3a. Leaves silvery pubescent abaxially ...............................................................................................................
    [Show full text]
  • Trillium Reliquum)
    REPRODUCTIVE BIOLOGY OF RELICT TRILLIUM (Trillium reliquum) Except where reference is made to the work of others, the work described in this thesis is my own or was done in collaboration with my advisory committee. This thesis does not include proprietary or classified information. _________________________________________ Melissa Gwynne Brooks Waddell Certificate of Approval: ________________________ _________________________ Robert Boyd Debbie R. Folkerts, Chair Professor Assistant Professor Biological Sciences Biological Sciences _____________________ _________________________ Robert Lishak Stephen L. McFarland Associate Professor Acting Dean Biological Sciences Graduate School REPRODUCTIVE BIOLOGY OF RELICT TRILLIUM (Trillium reliquum) Melissa Gwynne Brooks Waddell A Thesis Submitted to the Graduate Faculty of Auburn University in Partial Fulfillment of the Requirements for the Degree of Master of Science Auburn, Alabama August 7, 2006 REPRODUCTIVE BIOLOGY OF RELICT TRILLIUM (Trillium reliquum) Melissa Gwynne Brooks Waddell Permission is granted to Auburn University to make copies of this thesis at its discretion, upon request of individuals or institutions and at their expense. The author reserves all publication rights. ______________________________ Signature of Author ______________________________ Date of Graduation iii VITA Melissa Gwynne (Brooks) Waddell, daughter of Robert and Elaine Brooks, graduated from the University of North Alabama in 1996 with a bachelor’s degree in Geography and a minor in Biology. She graduated from Auburn University in 1998, in Horticulture and Landscape Design, and returned to Auburn University to pursue a master’s of science in 1999. Married in May 2004 to Erik Waddell, she accepted a position teaching seventh grade science and environmental science in December 2005. In July 2006, she begins a master’s degree in Education at the University of North Alabama.
    [Show full text]